Woodland Park Zoo’s only male lion was euthanized Wednesday due to kidney failure.

Xerxes, a 14-year-old South African lion who fathered three cubs, was known for his magnificent mane and his charismatic rumblings, according to the zoo’s blog.

“Xerxes roared like other lions do, but he also vocalized with a lot of low grumbles, high-pitched moans and a chuffing sound like a tiger,” said Martin Ramirez, mammal curator at Woodland Park Zoo. “He was very talkative with his lion keepers.”

He arrived at Woodland Park Zoo eight years ago under a breeding recommendation by the African Lion Species Survival Plan and fathered three cubs in 2014, all of whom have since moved to other zoos.

He was, according to zoo staff, an “excellent father … playing with and patiently tolerating the cubs’ rambunctious antics,” Ramirez said.

Xerxes was considered a geriatric lion, within the range of median life expectancy for male lions in zoos, according to the blog. He had been undergoing treatment for possible kidney disease. An exam scheduled for next week was moved up, with signs of decline and possible dehydration as Xerxes’ activity and appetite recently became low.

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While awaiting test results, signs of kidney disease became more apparent, according to Dr. Rob Browning, the park’s associate veterinarian. Xerxes showed initial signs of improvement, but another exam Wednesday indicated end-stage kidney failure.

“The death of Xerxes has hit our zoo family very hard,” Ramirez said.

As a standard procedure, the zoo’s animal health team will perform a postmortem exam to further diagnose factors that may have contributed to Xerxes’ decline and to share the results nationally among colleague scientists to help advance the understanding of medical issues in African lions.

In 2015, two African lion subspecies were listed under the Endangered Species Act, with a 40% decline in the wild population over the past two decades, according to the zoo. As few as 20,000 to 23,000 African lions are estimated to remain in the wild.